UDC 572

Human population physiology is a branch of physiological anthropology that studies the physiological structure of populations under specific environmental conditions. The theoretical basis of the direction is the concept of physiological homeostasis, and the methodological basis is consistency and integrity. The first task of human population physiology is to study the intra-and inter-group variability of the levels of individual physiological indicators and physiological status in general. A systematic approach allows us to establish differences or similarities in the physiological status of populations and explain them from the standpoint of ecological originality, state of adaptation and maladaptation.

Key words: physiological homeostasis, adaptation, environmental factors, physiological status, systematic approach, variability.

Introduction

In 1964, at the VII International Congress of Anthropological and Ethnographic Sciences (ICAEN), physiological anthropology was officially recognized as an independent scientific discipline. The founder of this discipline in Russian science can rightly be called Tatyana Ivanovna Alekseeva.

The content of physiological anthropology was defined as the study of adaptive variability - "a necessary prerequisite for the pan-ecumenical existence of a person" [Alekseeva, 1977, p. 6]. From a methodological point of view, an integrated approach to the study of various population groups living in various geographical conditions has become fundamentally new. A number of new features and methods were introduced into scientific use. A broad program of comprehensive research included the study of racial and diagnostic traits, morphological and physiological features of ethno-territorial groups, their genetic and demographic structure, the collection of nutrition data, and the determination of dermatoglyphic and odontological traits. In 1961, a team of employees of the Research Institute and the Museum of Anthropology of Moscow State University, led by T. And Alekseeva, started implementing this program. At the first stage, specific tasks were solved: the choice of characteristics and methods of their research; the study of their gender, age, and geographical variability. As a result, the geographical variability of the average values of the studied traits was found, which was interpreted in accordance with the scientific trends of those years: endogenous and exogenous determination.

Since the 1970s, the International Biological Program (IBP) has been conducting systematic research on human adaptation, primarily to extreme living conditions. Under the guidance of T. I. Alekseeva, the indigenous population of the Arctic zone was surveyed, and under the guidance of her student O. M. Pavlovsky, Aridna. In this decade, an ecological orientation has emerged in physiological anthropology and the work on complex programs has significantly expanded. Under the leadership of T. I. Alekseeva, a long-term study of the morphophysiological features of the indigenous population of the Altai-Sayan Mountains began-

as part of the Soviet-Mongolian expedition - the population of Mongolia (1976-1991).

During its 30-year existence (1961-1991), physiological anthropology was constantly enriched with new features and methods, the number of researchers dealing with the problems of this scientific discipline increased, and, naturally, its goals and objectives changed over time. The development of priorities: geographical variability-adaptation - ecology [Alekseeva, 1977, 1986, 1998]; the accumulation of results and the desire for their in-depth interpretation in separate sections of complex programs led to the natural emergence of a number of scientific directions within the framework of physiological anthropology.

Before proceeding to the description of human population physiology, it should be noted that this article does not mention numerous fundamental studies conducted in physiological anthropology by other domestic scientific schools, for example, the schools of MSU professors E. N. Chrysanfova and N. N. Miklashevskaya. This is due to the subject matter of the article: research on blood physiology was started under the guidance of T. I. Alekseeva and continued exclusively by her and her students. The author belongs to the school of T. I. Alekseeva and worked under her guidance from 1961 to 1972.*

Subject and objectives of human population physiology

From the very beginning of the formation of Russian physiological anthropology, the comprehensive program included the study of the levels of physiological parameters of blood. In the initial period, we mainly had to focus on foreign researchers: they were ahead of us in methodological terms, and they were able to collect a lot of factual material on the geographical variability of these indicators. The task of studying the intergroup variability of the average values of traits was to determine the so-called physiological norms characteristic of practically healthy people. It turned out that the "norms" vary significantly. When trying to explain this diversity, we took into account the main genetic factors and the influence of the environment, which, along with natural factors, includes the economic and social situation of the population, which determines the quantity and quality of food and energy expenditure associated with the specifics of labor. A review of the literature on the topic of interest to us has shown that during the half-century history of studying the levels of physiological blood parameters, there has been no progress in interpreting the ethno-territorial variability of their average values [Gudkova, 2008]. Therefore, the search for a single concept that can solve the problem of interpreting the results obtained by many researchers became an urgent necessity by the end of the XX century.

The causes and mechanisms of the observed phenomena were explained by the concept of physiological homeostasis (Cannon, 1932). Being dependent on environmental factors, it determines the diversity of the physiological structure of populations located in different environmental conditions and at different stages of adaptation. In human population physiology, homeostasis, adaptation, and environmental factors are discussed at the population level in terms of their causal relationship.

The methodological basis of the direction is consistency and integrity. Within the framework of systemic views, homeostasis is defined as the ability of a population to maintain dynamic stability of physiological status under changing environmental conditions, which is understood as a set of physiological variables that are interrelated at the organizational level and correlated at the population level. According to these definitions, the physiological structure of a population is perceived as an integral formation, which is provided by stable connections of individual parts of the general system. Today, there are many ways to assess these population characteristics (Gudkova, 2008). The biological meaning of the assessment is directly related to the viability of the population, which depends on the physiological homeostasis of the entire population, which consists of the physiological homeostasis of individual individuals.

The set of variables that make up the physiological status may vary depending on the goals and objectives of the study. The physiological parameters of blood we study (hemoglobin, whey proteins, total cholesterol, glucose) characterize individual metabolic properties of the body and take part in various energy processes. The levels of these indicators are quantitative phenotypic traits that vary within their reaction rate. On the one hand, the physiological characteristics of blood refer to "hard" constants, for which even a small change is a signal for the mobilization of homeostatic regulation systems (Anokhin, 1970). But, on the other hand, it is precisely because of their reactivity that they are convenient for studying adaptation, since, first of all, physiological reactions " underlie all adaptations-

* The author considers it his duty to express his deepest gratitude to Tatyana Ivanovna , a teacher and bright person who left so unexpectedly.

and, secondly, the degree of adaptability of a trait can be judged only in relation to its other states [Solbrig O., Solbrig D., 1982]. Therefore, extreme impacts are considered in human population physiology as a kind of model that reveals the features of physiological processes that are not manifested in normal conditions. Any extreme impact on the population stereotypically causes an increase in phenotypic variability due to a whole complex of genetic and physiological adaptation mechanisms (Sapunov, 1990).

The concept of variability is one of the basic concepts in human population physiology. With changes in the environment, the individual adaptability of organisms is of exceptional importance. At the population level, it is primarily expressed by an increase in the variability of physiological responses. The successful survival of the population as the main evolving unit within the species depends on its reserve. Only the presence of such variability and its maintenance from generation to generation within certain limits can protect a population from demographic catastrophes. Therefore, the main line of the new direction of physiological anthropology is the study of intra-group variability in the levels of physiological indicators in natural populations that live in rather contrasting conditions for a long time. And it is precisely for this reason that human population physiology is radically different from the traditional works of physiological anthropology, where the main criterion for successful adaptation is the average value of a particular trait. Such a statement of the question does not in any way negate the information value of the results obtained when comparing the average values of traits, but intra-population variability and its dynamics in stressful situations for a population will be a more accurate indicator of its adaptive capabilities.

Before proceeding to concrete examples, it makes sense to outline one more of the basic principles of human population physiology. The study of individual physiological traits as an end in itself, from the point of view of human population physiology, is not relevant, since the effect of population factors affects not individual traits, but their complexes [Zhivotovsky, 1991]. Analysis of each physiological variable as a component of the system is necessary, but this is only the first, although very important, stage of research. More detailed information is provided by studying the physiological status of the population as a whole. Using the methods of multidimensional biometrics, it is possible to assess the degree of similarity or difference in the physiological statuses of individual populations and understand the causes of intra-group variability in their physiological structure. For example, a logical analysis of the results obtained using the factor method using molecular biology data allows us to study physiological homeostasis at different levels of organization (from molecular to population) and determine internal and external parameters that affect its stability and variability [Gudkova, 2008].

In this article, two aspects of human population physiology will be briefly considered: the variability of physiological variables and the comparison of physiological statuses of populations as a whole structure.

Materials and methods

The relevance of comparing ecologically contrasting populations determined the choice of regions. Kamchatka and Khakassia are located in the temperate zone, Turkmenistan, Karakalpakstan, Kazakhstan - in the arid zone, and Chukotka - in the Arctic zone. The climatogeographic characteristics of the last two zones relate to extreme factors that cause a state of "dynamic mismatch" in the human body (Medvedev, 1979), which primarily leads to changes in physiological reactions. Most often, extreme weather is considered as a complex phenomenon that has a natural and anthropogenic origin.

The natural extremity of the arid zone is weaker than that of the Arctic zone. But in Central Asia, the seasonality of the climate is pronounced: from mid-May to mid-October-hot and dry, the rest of the year - wet and cold. This makes it possible to compare the physiological status of the same population under different conditions.

Unlike the indigenous desert population, people living in the Far North are constantly exposed to extreme factors of the Arctic zone, to which the Arctic aborigines have adapted for thousands of years. However, in recent decades, there has been a dramatic change in their way of life, and the indigenous population of the North is forced to adapt to a new heterogeneous environment, very different from that where most of its natural history has passed. In this regard, the aborigines of Chukotka experience physiological stress from a whole range of environmental factors, both climatic and geographical, and socio-cultural [Gudkova, 2008].

So, we studied samples from populations living in the temperate (Khakas, Kamchadals and migrants of Kamchatka), arid (Turkmens, Karakalpaks, Kazakhs) and Arctic (coastal Chukchi, Eskimos)regions zones. Bu-

Only men between the ages of 20 and 50 will be discussed; the sample size is 50 to 150 observations.

The levels of physiological blood parameters-hemoglobin, serum proteins (total protein, albumin, alpha1, alpha2-, beta -, gamma-globulins), total cholesterol-were determined by standard spectrophotometric methods. To correctly interpret their intra-group and inter-group variability, it is extremely important to observe methodological comparability.

To compare the physiological statuses of populations, the article uses one of the multidimensional statistical methods used in human population physiology - discriminant analysis. At the same time, the total protein level is excluded from the set of features as the sum of several variables (albumin and globulin levels), since such a variable does not carry any new information other than that contained in its components, and therefore is superfluous [Klekka, 1989]. The Statistica-6.0 program was used for calculations and graphical constructions.

Comparative analysis of intra-group variability of physiological traits

The integrating effect of a complex of environmental factors on a population depends on the characteristics of each individual and on the physiological differences between them. Adaptation to the external conditions of individual individuals, rather than groups or genes, culture or society (Foley, 1990), leads to the fact that in an extreme situation, a homeostatic disorder in the body of each person destabilizes the population-environment equilibrium system. In a changing environment, the population "should have a sufficient hereditary reserve of variability, which would create an opportunity for adaptive changes" [Levontin, 1981, p.246].

In the arid zone, during the hot season, under the influence of uncomfortable influences (high air temperature, defoliation), a disorder occurs in the homeostatic system of each person, causing, as it were, an expansion of the range of tolerance of the population [Gudkova, 2008]. Differences between the degrees of variability of adaptively significant features in the arid zone-hemoglobin levels (F - criterion - 2.46) and albumin levels (2.74) - in Turkmens examined in spring and autumn are significant with a probability of 0.99. An increase in intra-group variability and a larger range of variation series of the studied traits can be considered as a response of the population to signals of distress coming from the environment. In the" autumn " sample of Turkmens (examined in the hot season), the distribution of gamma-globulin levels (a sign reflecting the immune resistance of the population) is characterized by peaking, which may indicate a serious strain on the immune system in extreme conditions of the arid zone. In the" autumn "sample, compared with the" spring " sample (Turkmens surveyed before the onset of heat), there is a statistically significant asymmetry, indicating the predominance of individuals whose physiological homeostasis tends to maintain its stability, despite the impact of unfavorable climatic factors (Table 1). Thus, the different expression of fitness in individual individuals determines the population structure. variability of physiological responses, which, as noted above, are the basis of all adaptations.

In populations of the arid zone, without the influence of disturbing environmental factors, the variability of the levels of physiological blood parameters decreases, and the average values of the variables are very similar. This result can be considered as a manifestation of the stability of the levels of physiological blood parameters in the surveyed population.

The situation is different in Arctic populations (Gudkova, 2008). The average levels of physiological blood parameters in coastal Chukchi and Eskimos, despite the unity of environmental factors (socio-cultural and climatogeographic), differ significantly. Only the average values of albumin and gamma globulins were similar, but the intra-group variability of these traits was noticeably greater in Eskimos than in Chukchi. For example, the coefficient of variation in the level of gamma globulins is 41.4 and 26.7%, respectively. Variance analysis showed that differences between the degrees of variability of this trait (1.44) in Eskimos and coastal Chukchi are significant with a probability of 0.95.

When analyzing the distribution curves (tab. 2) for the levels of alpha2-, beta - and gamma-globulins in Eskimos, as well as for gamma-globulins and cholesterol in Chukchi, statistically significant positive (right-sided) asymmetry was obtained, which indicates the predominance of individuals with large and maximum values of traits in the samples. A statistically significant positive kurtosis that characterizes the island-vertex distribution, in turn, indicates a denser concentration of these features near the central trend. The results obtained indicate a violation of the physiological homeostasis of both populations, but the degree of response to stressors in coastal Chukchi and Eskimos is different. Although both groups are in extreme conditions, it seems that the Eskimos are in significant danger.

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Table 1. Statistical indicators of physiological traits in Turkmens

Table 2. Statistical indicators of physiological traits in Chukchi and Eskimos*

* See note. go to Table 1.

to a greater extent than coastal Chukchi, they experience physiological stress.

Quantitative traits that "carry a significant paralytic component" are more susceptible to changes depending on socio-ecological factors than others. Recently, a "so-called secular trend has been observed among many populations, including an increase in the average values of certain anthropometric characteristics, a decrease in their variability, and normalization of distribution curves." This trend is associated "with social progress, which contributes to a decrease in the environmental component of the variance of a quantitative trait and the most complete implementation of the genotype in the phenotype" [Kurbatova, Botvinev, Altukhov, 1991, p. 1229]. When trying to extrapolate the tendency in the variability of anthropometric traits to physiological ones, we come across a diametrically opposite situation in the Eskimos. Approximately 10 years after our work in Chukotka, E. R. Boyko [1990], having determined the metabolic parameters of only 16 Eskimos, obtained an average cholesterol level of 6.17 mmol/l, i.e., lower than in our sample (Table 2). Such a decrease can be considered a positive phenomenon, but it is very high for many Eskimos. the coefficient of variation (47.3 %) is discouraging for this trait. In the sample of Eskimos surveyed, it is equal to 17.0 %. As for the Chukchi people, the data presented in the work of E. R. Boyko coincide with our data (Gudkova, 2008).

So, physiological signs can vary within certain limits, corresponding to their genotype. To date, the assessment of the fitness of populations only by the average values of physiological variables "can be considered essentially primitive". Significantly more progressive is the assessment, which is based "on the analysis of a specific situation and a specific relationship, in which a positive result can be extracted, including from phenomena and relationships that are primarily perceived as destabilizing from the point of view of stability and norms" [Puzachenko, 1992, p.4]. Sometimes, only one form of statistical distribution of an adaptively significant trait can be used to assess the degree of population burden (Altukhov and Kurbatova, 1990). In this context, the intra-group variability of physiological traits is considered to be the main criterion for the state of adaptation of populations; and its analysis in dynamics will be the most prognostically reliable, which will allow us to separate temporary reversible changes (populations of the arid zone) of physiological homeostasis in the population-environment system from possible maladaptation (populations of the Arctic zone).

Discriminant analysis of intergroup differences in physiological status

Comparison of samples from populations in the temperate zone showed that the physiological status of Khakass people significantly differs from that of Kamchatka and Kamchadal migrants (Table 3, Fig. 1), which as groups from the same region without extreme environmental manifestations are close to each other (the differences are statistically unreliable: F= 1.55; p < 0.15). The albumin level has the highest standardized coefficient in the samples of the temperate zone. All other signs are at least twice as significant. It should be noted that the contribution of variables to the value of discriminant functions in the analysis of the physiological status of male samples from temperate zone populations is logical and corresponds to the ideas of sexual dimorphism in the levels of physiological indicators

Table 3. Discriminant analysis results

1. Discriminant analysis of the physiological status of samples from temperate zone populations.

1-Kamchadals; 2-migrants of Kamchatka; 3-Khakas.

Figure 2. Discriminant analysis of physiological status

samples from arid zone populations.

1-Turkmens surveyed in autumn; 2-Karakalpaks; 3-Kazakhs; 4-Turkmens surveyed in spring.

See Table 4. Classification matrix of samples from temperate zone populations (percentage)

blood pressure (the maximum load in the female samples falls on the level of gamma globulins).

During the classification procedure, it turned out that the largest percentage of individuals correctly classified according to their physiological status was in the sample of migrants (Table 4). The percentage content as a measure of forecast accuracy refers to the most important discriminant information. The proportion of correct predictions in the Kamchadal sample cannot be considered correct, since the expected accuracy of random classification of three samples is 33 %. The relatively high percentage of Kamchadals diagnosed as migrants indicates, in turn, that the physiological status of these two populations is similar. Perhaps, in addition to the ecological factor, the result was also influenced by the mixed origin of the Kamchadals. Thus, it is not possible to identify Kamchadals by their physiological status with sufficient accuracy.

Analysis of materials from the arid zone showed statistically significant differences in the physiological status of all samples (see Table 3, Figure 2). The peculiarity of the Kazakh sample was once again revealed (Gudkova, 2008); its "transitional" essence at the time of the survey: a large intra-group variability, as in the "autumn" sample of Turkmens, and convergence with the "spring" sample in centroids in the coordinates of the first discriminant function, where the maximum significance of canonical variables determines the level of albumin. This trait, which plays an important role in adaptation to extreme factors of the arid zone, is also most significant in female samples. In the second discriminant function, the hemoglobin level has the highest standardized coefficient (the same in women), which is also an adaptively significant feature in hot desert climates. Thus, extreme pressure clearly separated the physiological status of Turkmens and Karakalpaks surveyed in the hot season from that of Turkmens from the "spring" sample and Kazakhs by the first discriminant function. Anthropogenic factors (cotton defoliation) that affected the physiological status of Karakalpaks determined, in turn, the separation of the system of variables between them and Turkmens.

In the classification of samples from populations of the arid zone, discrimination between two samples of Turkmens surveyed in different seasons is of the greatest interest (Table 5). It turned out that only 2.5% of Turkmens from the "autumn" sample fall into the "spring" sample, and none of the latter is diagnosed as being surveyed in autumn. Percentage of correct classifications (for four

See Table 5. Classification matrix of samples from arid zone populations (percentage ratio)

The random variant (25%) can be considered significant; the proportion of accurate predictions is especially high in the sample of Turkmens surveyed in an extreme situation. Interestingly, despite the impact of anthropogenic factors affecting the levels of individual physiological blood parameters, the physiological status of Karakalpaks in general turned out to be quite similar to that of Turkmens from the "autumn" sample: 30% of Karakalpak men were included in this sample.

Populations of the Arctic zone are represented in our study only by Eskimos and coastal Chukchi, and to obtain a two-dimensional graphic image under the conditions of discriminant analysis, it is necessary to operate with at least three populations, so a sample from the temperate zone was taken. According to the traditions established in anthropology, and due to the absence of an ecologically extreme situation in this zone, the most suitable background for analyzing the differences in the physiological status of Eskimos and Chukchi is a sample of migrants from Kamchatka.

Discriminant analysis of the system of physiological features in the graphic image (Fig. 3) gives a generalized idea of the intermediate position of the Eskimos between migrants and Chukchi. All differences are statistically significant (see Table 3). The obtained result is well interpreted from the point of view of physiological homeostasis disorders in Eskimos, for example, under the influence of the transition to the "European style" of nutrition. The same conclusion can be reached by looking at standardized coefficients. The highest loads for the first discriminator fall on the levels of beta-globulins. The maximum contribution to the value of the second function is made by the levels of alpha2-globulins. Note that 35 % of the variability that it describes is quite large. Therefore, the most significant loads on both functions on the levels of beta-and alpha2-globulins-signs associated with carbohydrate and lipid metabolism-are of some importance for interpreting the physiological status of Chukotka aborigines [Gudkova, 2008].

The classification procedure showed (tab. 6) that the proportion of Eskimos correctly classified according to their physiological status is significantly lower than that of coastal Chukchi. The discriminant information provided by the classification matrix clarified the position of the Eskimos shown in Fig. 3. According to their physiological status, 16 % of Eskimos are included in the sample of Chukchi, 22% - in the sample of migrants. As can be seen, the classification procedure for the Eskimo population was not effective enough:

3. Discriminant analysis of the physiological status of samples from populations of the Arctic zone.

1-Eskimos; 2-coastal Chukchi; 3-migrants of Kamchatka.

Table 6. Classification matrix of samples from Arctic zone populations (percentage ratio)

the percentage of correct definitions is significantly lower than in the Chukchi and migrant samples. Thus, the classification results reflected the peculiarities of the physiological status of Eskimos.

Thus, the value of the canonical correlation, the values of the Wilks l-statistics and standardized coefficients, the classification matrix, and, finally, the ability to visually perceive the division of samples according to the system of characteristics - all this makes discriminant analysis an extremely sensitive tool for assessing inter-group differences in the integral structure of the physiological status of populations.

Conclusion

Traditional methods for studying the physiological status of populations include determining the arithmetic mean, variance, coefficient of variation, and analyzing the distribution curves of individual physiological traits. In the system approach, correlation is considered as one of the essential integration mechanisms, and therefore the correlation method is also widely used in human population physiology. The relevance of studying the physiological status as an integral structure was determined by the use of multidimensional biometrics methods (factorial, discriminant, cluster, and canonical). They provided new opportunities for analyzing the intra-and inter-group variability of a system of variables. Multidimensional methods allow us to establish differences or similarities in the physiological status of populations and explain them from the standpoint of ecological originality, adaptability or maladaptation.

Various assessments of the physiological state of a population can be successfully applied when considering environmental problems of the population. For example, you can calculate an average indicator of the variability of physiological variables as components of the system to see the adequacy of the population to its environment. The impact of a number of ecologically significant anthropogenic factors causes significant changes in the physiological homeostasis of populations. The correct interpretation of these changes using various biometric methods is essential for the development of effective measures to protect the population.

List of literature

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Altukhov Yu. I., Kurbatova O. L. Problems of adaptive norm in human populations // Genetics. 1990, Vol. 26, No. 4, pp. 583-598.

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The article was submitted to the Editorial Board on 15.05.08.

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